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The hierarchical nature and processes of different levels of ecological systems: Species Interactions Ch 17 Herbivory Objectives • Herbivory Effects on plants How demonstrate herbivory effects Herbivore selectivity Plant deterrents to herbivory What limits herbivory? • Pairwise interspecific interactions ***Which is: +/-, +/0, +/+, -/-, -/0 ? • • • • • • • • Mutualism Commensalism +,0 Amensalism -,0 Herbivory Predation Parasitism Disease Competition Pairwise interspecific interactions • • • • • • • • Mutualism (+/+) Commensalism (+, 0) Amensalism (-, 0) Herbivory (+/-) Predation (+/-) Parasitism Disease Competition (-/-) Fluidity of interspecific relationships: • Can evolve from one type to another. • Switch + and - signs of interaction, e.g. +/+ to +/-. In food chains, all life forms are both consumers and victims of consumers. • • • • • Predators Parasites Parasitoids Pathogens Herbivores Herbivory: Effects on plants Outbreaks of herbivorous insects can defoliate forests. Spruce budworm Herbivory has great effects on plants: 1) individual/ecosystem 2) population 3) community • If prefer dominant species---> • What is a keystone • herbivore? • If prefer subdominant species---> Figure 1 How do we test the hypothesis that herbivores control plant populations? Describe, then explain these results. Herbivores feed on plants - and also inoculate them with pathogens and rot-causing microbes. Does herbivore control plant species? If…. then… • Natural enemies hypothesis: • Biological control: Figure 2 In spite of plant defenses, herbivores can control plant population size. Klamath weed + beetle (biological control agent) ***What are plant deterrents to herbivory? • Structural defense • Low nutritional content; sequester nutritious parts • Mutualistic defense (ant-acacia) • Secondary compounds Figure 3 Chemical defenses (secondary compounds) are toxic to herbivores. Types of secondary compounds • growth regulators • toxins against generalist herbivores specialists evolve to detoxify toxin often N-based lignin, alkaloids, non-protein amino acids, cyanogenic glycosides--->HCN • digestive inhibitors against specialists often C-based tannins, phenolics, terpenoids Chemical defenses • Constitutive: high levels at all times • Induced: increase greatly after • attack • Theory: Cost of defense is too high to maintain under light herbivory. • (but how quickly can they make them? • Hypothesis: Plants ‘eavesdrop’ on neighbors signal to make defense. What is evidence that plant defenses are induced by herbivory? #2 to #1 • Mite sp 1 attacks. • Plant responds by making defense chemical. • Mite sp 2 attacks but in much lower numbers. ***Are herbivores ‘lawnmowers’ or selective feeders? What 3 factors may explain results? Figure 4 ***Describe the major pattern in this figure. Generate a WHY ? Develop an ‘If…then’. Herbivores of oak leaves April October Figure 5 Hypothesis/prediction: • If oak leaves become less suitable insect food as they age, • then caterpillars fed young leaves will grow better than if fed slightly older leaves. • Diet larval weight • young leaves 45 • old leaves 18 % adults emerge 76 0 • ***What is conclusion? • Do data support the hypothesis? “Figure” 6 ***What are three changes as a leaf ages that could account for the previous results? (3 alternative hypotheses) • H1: Increase in toughness • H2: Increase in secondary chemicals • H3: Decrease in nutrient quality Toughness index Figure 7 Leaf age ***Develop predictions for H1 toughness and H2 chemical defense. • If leaf toughness explains seasonal feeding pattern of oak insects, • then larvae should grow equally well when eating ground-up old vs. young leaves. • If chemical defenses have increased with leaf age, • then larvae should grow better on ground-up leaves of young than old leaves. • ***Are predictions ‘operational’? • Do they contain independent and dependent variables? ‘Figure’ 8 ***Results • Larvae fed ground-up leaves • Young leaves • Old leaves Larval weight 37 35 • ***Which hypothesis is supported? • Why hasn’t NS favored insect mouth parts able to cope with tough leaves? • 3rd alternative hypothesis is still possible; Maybe poorer nutrition in later summer; then NS toward early feeding. “Figure” 9 • Herbivores consume only @ 10% of plant productivity (up to 30-60% in grasslands). Why so little? • • • • • • ***What factors limit herbivory? Predators Herbivores Plants Nutrients Abiotic factors • The ‘world is green’ hypothesis: Herbivores consume a small % of vegetation because they are held in check by a variety of factors. Top-down control predators Tri-trophic interactions herbivores plants nutrients/light Bottom-up control “Figure” 10 Observation/question: Despite many potential herbivores, why do leaves lose low leaf area? • Observation: Birds eat insect herbivores. • ***Hypothesis: • If bird predation on insect herbivores indirectly reduces the amount of leaf area consumed, • ***Prediction: then leaf area consumed will be greater for plants with bird-exclusion cages than those without cages. ‘Figure ‘ 11 Experimental set-up…caged tree saplings “Figure” 12 1) number of insects: 70% greater on saplings without birds than with birds. 2) % leaf area missing: 35% without birds 22% with birds Results: • *** What’s conclusion? • Support for hypothesis? • Tri-trophic interaction; top-down control. • Bird predation: directly reduces # of herbivores indirectly reduces leaf damage by herbivores • New questions: Will 1) decreases in bird populations due to forest fragmentation or 2) change in phenology increase insect damage?